DEPARTEMEN KEHUTANAN REINWARDTIA Vol 12, Part 5, pp: 391 – 404 391 FLORISTIC COMPOSITION AND STRUCTURE OF SUBALPINE SUMMIT HABITATS ON MT. GEDE-PANGRANGO COMPLEX, CIBODAS BIOSPHERE RESERVE, WEST JAVA, INDONESIA Received December 1, 2008; accepted December 3, 2008 A. SADILI Herbarium Bogoriense, Botany Division, Research Center for Biology – LIPI, Jl. Raya Jakarta Bogor Km 46, Cibinong 16911, Indonesia K. KARTAWINATA Herbarium Bogoriense, Botany Division, Research Center for Biology – LIPI, Jl. Raya Jakarta Bogor Km 46, Cibinong 16911, Indonesia; UNESCO, Jakarta Office, Jakarta, Indonesia. E-mail: kkjak@indo.net.id (author for correspondence) A. KARTONEGORO Herbarium Bogoriense, Botany Division, Research Center for Biology – LIPI, Jl. Raya Jakarta Bogor Km 46, Cibinong 16911, Indonesia H. SOEDJITO Herbarium Bogoriense, Botany Division, Research Center for Biology – LIPI, Jl. Raya Jakarta Bogor Km 46, Cibinong 16911, Indonesia; Indonesian National MAB Committee, LIPI, Jakarta, Indonesia A. SUMADIJAYA Herbarium Bogoriense, Botany Division, Research Center for Biology – LIPI, Jl. Raya Jakarta Bogor Km 46, Cibinong 16911, Indonesia ABSTRACT SADILI, A., KARTAWINATA, K., KARTONEGORO, A., SOEDJITO, H. & SUMADIJAYA, A. 2009. Structure and composition of subalpine summit habitats on Mt. Gede-Pangrango complex, Cibodas Biosphere Reserve, West Java, Indonesia. Reinwardtia 12 (5): 391–404. — We undertook a phytosociological analysis of the subalpine herbaceous and shrubby vegetation at the Mandalawangi and Suryakencana meadows and the scrub at the Crater Side at the tops of Mt. Gede and Mt. Pangrango in the Cibodas Biosphere Reserve. We recorded 30 species of 18 families of saplings, shrubs, seedlings and herbs in 78 quadrats with a total area of 7,800 m2. Anaphalis javanica, a woody tall herb and long-lived pioneer was the dominant species in the sapling and shrub stratum, while Isachne pangerangensis, Tripogon exiguus and Carex verticillata were prevalent in the seedling and herb stratum at Mandalawangi and Suryakencana. Stunted shrub is Vaccinium varingaeifolium, dominant in the Crater Side scrub. Based on the importance values, the Mandalawangi meadow may be designated as the Anaphalis javanica-Isachne pangerangensis community type, the Suryakencana meadow as Anaphalis javanica-Tripogon exiguus community type and the Crater Side scrub as Vaccinium varingiaefolium-Seliguea feei community type. The similarity indices between Mandalawangi and Suryakencana community types were very high (>75 %) while those between the Crater Side and Mandalawangi and the Crater Side and Suryakencana were very low (<10 %). Poor soil conditions and fire seem responsible for the perpetual existence of A. javanica. Keywords: Structure and composition, subalpine meadows, Anaphalis javanica, top of Mt Gede-Pangrango ABSTRAK SADILI, A., KARTAWINATA, K., KARTONEGORO, A., SOEDJITO, H. & SUMADIJAYA, A. 2009. Struktur dan komposisi habitat subalpin di puncak G. Gede-Pangrango, Cagar Biosfer Cibodas, Jawa Barat, Indonesia. Reinwardtia 12 (5): 391–404. — Analisis fitososiologi telah dilakukan pada vegetasi terna di alun-alun Mandalawangi dan Suryakencana dan semak di Tepi Kawah daerah puncak G. Gede dan G. Pangrango di Cagar Biosfer Cibodas. Tercatat sebanyak 30 spesies dari 18 suku belta, perdu, semai, dan terna dalam 78 petak cuplikan dengan luas total 7.800 m2. Anaphalis javanica, terna tinggi mengayu dan pionir berumur panjang, adalah spesies dominan pada stratum belta dan perdu , sedangkan Isachne pangerangensis, Tripogon exiguus dan Carex verticillata adalah jenis utama pada stratum semai dan terna di alun-alun Mandalawangi dan Suryakencana. Perdu kerdil Vaccinium varingaeifolium adalah jenis dominan dalam komunitas semak Tepi Kawah. Berdasarkan nilai pentingnya vegetasi di alun-alun Mandalawangi dapat disebut sebagai tipe komunitas Anaphalis javanica-Isachne pangerangensis, di alun-alun Suryakencana tipe komunitas Anaphalis javanica-Tripogon exiguous dan semak Tepi Kawah tipe komunitas Vaccinium varingiaefolium-Seliguea feei. Indeks kesamaan antara tipe komunitas di mailto:kkjak@indo.net.id REINWARDTIA [VOL.12 392 Mandalawangi dan di Suryakencana sangat tinggi (>75 %), sementara antara tipe komunitas Tepi Kawah dan Mandalawangi serta antara tipe komunitas Tepi Kawah dan Suryakencana sangat rendah (<10 %). Kondisi tanah miskin dan juga kebakaran tampaknya merupakan faktor yang mempertahankan kelanggengan A. Javanica. Kata kunci: Struktur dan komposisi, alun subalpin, Anaphalis javanica, puncak G. Gede-Pangrango INTRODUCTION The Cibodas Biosphere Reserve comprises natural ecosystems contained within the Gunung Gede-Pangrango National Park and man-made ecosystems surrounding the Park bordered by the encircling highway connecting Ciawi-Sukabumi- Cianjur-Puncak-Ciawi (Figure 1). The Cibodas Biosphere Reserve is listed as a member of the World Network of Biosphere Reserves (Rustiami, 2004). It is an important conservation and biological research site that has a long history in Indonesia. The natural ecosystems within the Park have a high plant species richness, where to date 823 species of flowering plants have been recorded (Sunarno & Rugayah, 1992) and the number have increased with additions from recent inventory and ecological studies. The site has attracted a large number of scientists to explore and study the tropical biology and ecology of plants and animals in these ecosystems since the establishment of the Botanic Gardens in Bogor in 1817. The brief history of scientific research in the area is included in the Mountain Flora of Java by Van Steenis et al. (1972, 2006), while earlier Van Steenis & Van Steenis-Kruseman (1953) listed botanists and their publications based on research in Mt. Gede-Pangrango up to 1952. All of them were the results of qualitative observations. Some of the noted earlier botanists and plant ecologists were Blume (1825), Docters van Leeuwen (1933), Junghuhn (1845), Hasskarl (1840), Koorders (1918–1923), Korthals (1848), Kramer (1926, 1933), Reinwardt (1819), Seifriz, (1923), Teysmann (1842), and Went (1940). After the 2nd World War research in the area has been undertaken by Abdulhadi et al. (1998), Meijer (1959), UNESCO et al. (1975), Srijanto (1987), Sunarno & Rugayah (1992), Yamada, (1975, 1976, 1977), and several undergraduate and graduate students from various universities. Quantitative ecological studies in the natural ecosystems of the Cibodas Biosphere Reserve have been concentrated on the montane and subalpine forests (Abdulhadi et al., 1998; Srijanto, 1987; UNESCO et al., 1975; Yamada, 1975, 1976, 1977); and none has been conducted on subalpine herbaceous and shrubby communities. The following account reports the results of a quan- titative phytosociological study in the alun-alun (open herbaceous and shrubby meadows) Mandalawangi and Suryakencana and the crater side scrub at the tops of Mt. Gede and Mt. Pangrango, complementing the qualitative records of vegetation made earlier by various botanists, particularly Docters Van Leeuwen (1933) and Van Steenis et al. (1972, 2006). STUDY SITE The Cibodas Biosphere Reserve comprises natural ecosystems contained within the Gunung Gede–Pangrango National Park (GGPNP) and man-made ecosystems surrounding the Park, bordered by the intercity highway connecting Ciawi-Sukabumi-Cianjur–Puncak–Ciawi, with the total area of 108,000 hectares. It is located within Bogor, Cianjur and Sukabumi Districts, West Java (Figure 1). The GGPNP comprises an area of 21,975 hectares and was established initially in 1980 through the Decrees of the Minister of Agriculture in 1980 and was extended in 2003 through the Decree of Minister of Forestry in 2003 (Balai TNGGP, 2007). The park area starts at an elevation of 700 m (Helmi et al., in preparation) and extends upwards to the areas of twin mountains, Mt. Gede (2,962 m alt.) and Mt Pangrango (3,019 m alt.), connected by a sadle at 2,400 m alt. at Kandang Badak. It covers upper lowland forests, montane forests, subalpine forests, subalpine scrubs, subalpine grasslands, subalpine herbaceous communities and subalpine swamps (for detailed descriptions see Docters Van Leeuwen, 1933; Van Steenis et al., 1972, 2006 a & b; Yamada, 1975, 1976, 1977). On the tops of Mt. Gede-Pangrango, the subalpine vegetation dominates the landscape and consists of open herbaceous and shrubby meadows locally known as the alun-alun dominated by Anaphalis javani- ca, scrubs dominated by Vaccinium varingiaefo- lium and low forests dominated by Leptospermum flavescens or Albizia lophanta. The climate diagram (Figure 1) shows the mean monthly rainfalls and temperature at the top of Mount Pangrango. The mean annual rainfall at the top of Pangrango is 3369 m and at Kandang Badak is 3818 mm (LMG, 1969). In the Schmidt & Ferguson (1951) scheme, the rainfall in the 2009] SADILI et al.: Floristic and structure of subalpine summit on Mt. Gede Pangrango 393 . Sukaraja SUKABUMI Selabintana Cimungkat Nagrak Bodogol Cimande Tapos Cisarua Gunung Mas Puncak BOGOR Ciawi Cisaat Karang Tengah Gadog Cimacan Cipanas Gn. Putri Sarongge Gedeh Gekbrong Cisarua Mt..Pangrango Situgunung Cibodas STUDY SITE N Mt..Gede CIANJUR Cicurug INDONESIA CIBODAS BIOSPHERE RESERVE (108.000 Ha.): Cibadak Core zone (15,196 Ha.): Conservation area covering montane forest, subalpine forest, and subalpine scrub and herbaceous vegetation. Buffer zone (12,700 Ha) : It comprises Cibodas Botanical Garden, Safari Garden, tea plantation, and planted forests. Transition zone (80,104 Ha): Development area comprising, rice fields, home gardens, dry land crop gardens, settlements, industries, recreation areas, etc. Figure 1. The Cibodas Biosphere Reserve bordered by the intercity highway connecting Ciawi, Cianjur and Suka- bumi (Redrawn after TNGP cited by Rustiami, 2004). The climate diagram shows the mean monthly rainfall and temperature at the top of Mount Pangrango (3019 m alt.) (Data from LMG 1969 & Docters Van Leeuwen, 1933). MT . P ANGRANGO ELEVAT ION : 3019 M ANN. T EMP .: 17.8O C ANN. RAINF.: 3369 MM 0 10 20 30 40 J F M A M J J A S O N D J TEMP. (0C) 0 20 40 60 80 RAINFALL (X 5 MM) MEAN T EMP . MEAN RAINF. REINWARDTIA [VOL.12 394 entire park, including the area of the tops of Mt. Gede–Pangrango, belongs to the Type A, which is very wet. The rainfall is not uniformly distributed throughout the year, but there is a 2 month dry period in July and August, with the mean monthly rainfall of 80 and 85 mm, respectively. Data on the mean number of rain days shows that the dry season has little effects; during the driest month in July and August, where the number of rain days was 16.4 and 14.7, respectively (LMG, 1969) implying that rain falls every two days with a significant measurable amount. The monthly mean of daily air temperature at the top of Pangrango ranges from 8.4°C to 9.7°C with the annual mean of 9.0°C, the highest mean daily temperature varies from 11.3°C to 11.9°C between 10 AM. and 2 PM. (Docters Van Leeuwen, 1933). In the Gede–Pangrango mountain top area, in addition to rainfall, precipitation from the fog that develops almost every day and forms dew on leaves and mosses in the forest is significant also, but to date no data are available (Van Steenis et al., 1972, 2006) The soils in the top area consist mainly of stones and gravels deriving from lava and volcanic ash which may be designated as grey regosols and lithosols, while on the forested slopes all belong to the mixture of andosol and latosol or latosol and regosol (FAO, 1978) The study sites were located in the Alun-alun Mandalawangi and Alun-alun Suryakencana and a site of open low woody vegetation near the crater of Mt. Gede, which for short will be called Mandalawangi, Suryakencana and the Crater Side, respectively. Herbaceous vegetation covers the alun-alun and always looks dry, especially grasses and Anaphalis javanica. The alun-aluns are surrounded by subalpine forests, dominated jointly by Vaccinium varingiaefolium and Leptospermum flavescens or in places by Albizia lophanta. Hikers have frequently visited the sites and in places left disturbances, although as a whole the areas are still in natural state. The severe disturbance was attributed to fire, which apparently has frequently occurred in the surrounding forests. Docters Van Leeuwen (1933) reported fire occurred on the western side of Mt. Gede and in the present observation it affected the forests on the northern side of a smaller mountain, Mt. Gumuruh. In 2006, fire spread from an abandoned campfire and burnt part of the Anaphalis javanica stand in the western section of Suryakencana. The Alun-alun Mandalawangi (Figure 7) is located at 2,978 m alt. at 06° 46.403‘ E and 106° 58.042’ S, about 100 m north-east from the top of Mt Pangrango at 06° 46.225’ E and 106° 57.913’ S. The alun-alun is a meadow located on a basin derived from the inactive crater and now forms an open and roundish field with diameter of about 250 m, making a total area of about five hectare. The central section of the field is somewhat flat, consisting of sand and gravel, while the outer section towards the edge is sloping and covered by A. javanica, where the soil layer is of 3–5 cm thick. In the middle, a small river is running from East to West. The field is surrounded by forest, loaded with Usnea sp. and other mosses, with soil of about 10 cm thick. During the study on August 2007, the weather was dry with temperature measurements averaging 4°C at 6 AM., 18°C at Noon, and 12°C at 9 PM. Fog descended every day between 4 and 8 PM. The Alun-alun Suryakencana (Figure 8) is a sickle-shaped meadow, developed from the extinct volcanic crater of about 2 km long and about 200 m wide, stretching from north-east to south-west, making up a total area of 49.72 hectares. It is located about 200 m below the top of Mt. Gede on the narrow gap separating Mt. Gede on the north side and smaller mountain Mt. Gumuruh on the south side at 06° 46.200’ E and 106° 57.821’ S at an altitude of 2,719 m. On the west side the land is sloping toward the center where a relatively big stream is running, with smaller streams coming from the gentle slopes around it. During the rainy season the river is overflowing (Van Steenis et al., 1972, 2006), but during the dry season such as during the present study the river is dry. On the east side the slope is more gentle, especially from the middle section toward the eastern edge of the field. The soil in the central section of the field is yellow, thin and mixed with sand and gravel, while at the edge next to the forest the dark yellow soil is about 5–10 cm thick, mixed with gravels and small boulders or tallus, where stunted A, javanica grows and the entire soil surface in between A, javanica is covered by lichen Stereocaulon graminosum and mosses, especially Rhacomitrium lanuginosum. The air temperature measured during the study was 5°C (at 6 AM.), 22°C (at Noon) and 2°C (at 9 PM.). The Crater Side (Figure 9) is an area of about two hectares, which is located at 2,514 m altitude near two craters (Kawah Lanang and Kawah Wadon), at 06° 468.68’ E and 106° 58.863’ S. The site is sloping towards the east, ending at the edge of a very steep ravine, and the slope varies up to 40%. The substrate mainly consists of boulders of varying sizes up to 2 m in diameter. The vegetation is a sparse scrub, composed mainly of dwarf trees of Vaccinium varingiaefolium, Albizia lophanta and Leptospermum flavescens, as 2009] SADILI et al.: Floristic and structure of subalpine summit on Mt. Gede Pangrango 395 well as tall herb Anaphalis javanica, growing between boulders. The area is very dry where beard-moss Usnea and other species of mosses can hardly grow. METHOD The quadrat method was used to sample the Mandalawangi and Suryakencana meadows and the scrub on the Crater Side. Since there were no trees, the vegetation was stratified into two strata only: (1) Sapling and shrub stratum, where a sapling or shrub is defined as a woody plant with DBH (Diameter at Breast Height) ≤10 cm with height of ≥ 50 cm; tall herb is included here also, and (2) Seedling and herb stratum, where a seedling or a herb is defined as a plant with height of ≤50 cm. Twenty three (10 x 10 m) quadrats with the total area of 2,300 m2 were established and distributed systematically with intervals of 20 m in Mandalawangi, 50 quadrats with the total areas of 5,000 m2 in Suryakencana, and five quadrats with a total area of 500 m2 in the Crater Side. Each quadrat was divided into four (5 x 5 m) plots for recording saplings and shrubs and a 1 x 1 m subplot for seedlings and herbs, which was nested within one of the four plots within each quadrat. Different life forms were identified to the species level, their heights were measured and the percentage of cover of each species was estimated. The number of individuals of each species within the quadrats, plots and subplots were counted. Additional information on soil and substrate as well as other habitat conditions for each quadrat were noted and flowering status of species were recorded. Data were analyzed using the vegetation parameters of dominance (as expressed by the percentage of cover), density and frequency of each species, whose relative values were summed up and presented as the Importance Values, expressed in percentages (Mueller–Dombois & Ellenberg, 1974). Voucher specimens were collected and identified at the Herbarium Bogor- iense in Cibinong. The species nomenclature follows Flora of Cibodas (Sunarno & Rugayah, 1992) and Flora of Java (Backer & Bakhuizen van den Brink Jr., 1963-1968). RESULTS AND DISCUSSION A total of 30 species of 18 families of saplings, shrubs, seedlings and herbs were recorded in 78 quadrats in the study sites. In the sapling and shrub stratum only 18 species were recorded in the vegetation sampled in Manda- lawangi (13 spp.), Suryakencana (14 spp.) and the Crater Side (10 spp.). Of the total of 18 species of saplings, shrubs, and woody herbs, seven species were common to the three sites, four species in Mandalawangi and Suryakencana, and one species in Suryakencana and the Crater Side, while species with restricted distribution were only two in Mandalawangi, Suryakencana and the Crater Side, respectively (Annex 1). In the sapling, shrub and woody herb stratum, the Shannon diversity indices of sapling, shrub and woody herb communities in the three sites are low, i.e. 1.69 for Mandalawangi, 1.67 for Surya- kencana and 1.16 for the Crater Side. The species area curve (Figure 3) shows that the number of species increases rapidly up to 1,500 m2 and from this point onwards the increase is slight, confirming further the low species diversity. It may be implied also that the minimum area for this sapling community is 1,500 m2. In this stratum, it can be noted further that Anaphalis javanica was dominant, as indicated by its high importance value and relative dominance in Mandalawangi and Suryakencana, but its presence in the Crater Side was insignificant. (Figure 2, Annex 1) The next important species were Vaccinium varingiaefolium, Rhododendron retusum and Gaultheria punctata, which were dominant and co-dominant species in the Crater Side. They were less significant in Mandalawangi and Suryakencana, and even Rhododendron retusum was absent. Albizia lophantha and Leptospermum flavescens were the prominent tree species in these sites, but they were mostly crooked and low in stature, hence constituted only the component of the sapling and shrub stratum. They grew well at the edges of the alun-alun, merging with surrounding forests dominated by L. flavescens, which could reach the height of more than three meters. This is apparently the poorly developed variant of the subalpine forest occurring in the summit area described quantitatively by Yamada (1977) and earlier qualitatively by Docters Van Leeuwen (1933) and Van Steenis et al. (1972, 2006), where A. lophanta, L. flavescens and V. varingiaefolium were the leading species. A. javanica was widespread throughout, but formed dense stands at the edge of Mandalawangi and Suryakencana. In contrast, it did not develop well and was sparsely distributed in the Crater Side, where it grew only on firm and hard soils between boulders. Outside the quadrats it could be found also growing in the valleys of dead and active craters as well as on poor, sandy and stony soils. A. javanica groves developed densely on the REINWARDTIA [VOL.12 396 slope and formed a tall scrub with height up to 1.5 m. The soils were covered by mosses with various herbaceous species growing in between. Quanti- tatively there was no consistent correlation between percentage of cover and habitat (slope, soil depth and soil pH), but qualitatively it could be observed that A. javanica grew in roups on open and flat lands or gentle slopes with poor sandy or stony soils as well as volcanic rocks. Our observations confirm the earlier work by Docters Van Leeuwen (1933) and Van Steenis et al. (1972, 2006), who noted also that from time to time A. javanica groves died because of frosts that took place quite frequently at night. Docters Van Leeuwen (1933) noted that various aspects of alun-alun had not changed since they were described by Junghuhn (1845) and later by Korthals (1848). A. javanica occurred also in the alun-alun at the top of Mt. Papandayan, West Java (Van Steenis, 1932), and less abundantly at Mt. Ceremai, West Java (Lam, 1925), Mt. Sumbing and Mt. Sindoro in Central Java (Docters Van Leeuwen, 1930). The occurrence of apple trees, Malus domes- tica, in Mandalawangi and Suryakencana is interesting. They were the survivor of a variety of temperate fruit and vegetable species planted in an experimental garden at Kandangbadak by Teysmann in 1839, to cater the need of the Governor General for European vegetables and fruits (Van Steenis et al,. 1972, 2006). SAPLING & SHRUB 0 50 100 150 200 250 Scheflera lucescens Rhododendron retusum Albizia lophanta Dichrochepala chrysantemifolia Rubus sp. Hypericum leschenaultii Vaccinium laurifolium Eurya glabra Rubus lineatus Malus domestica Photinia integrifolia Lonicera javanica Symplocos cochinchinensis Rapanea avenis Gaultheria punctata Vaccinium varingiaefolium Leptospermum flavescens Anaphalis javanica IMPORTANCE VALUE (%) MANDALAWANGI SURYAKENCANA CRATER SIDE Figure 2. Importance Values of saplings and shrubs at Mandalawangi, Suryakencana and Crater Side on the tops of Mt. Gede and Mt. Pangrango, Cibodas Biosphere Reserve. 2009] SADILI et al.: Floristic and structure of subalpine summit on Mt. Gede Pangrango 397 In the seedling and herb stratum altogether 21 species were recorded, of which 11 species occurred in Mandalawangi, 16 species in Suryakencana and five species in the Crater Side (Figure 4, Annex 2). The Shannon diversity indices for the three communities in this stratum are also low, i.e., 1.82 for Mandalawangi, 2.17 for Suryakencana and 0.56 for the Crater Side. The species-area curve (Figure 5) shows that the number of species of seedlings and herbs increases rapidly up to 20 m2 and from this point onwards the increase is slight, confirming further the low species diversity. It suggests that the minimum a- rea for this seedling and herb community is 20 m2. Figure 3. Species-area curves of saplings, shrubs and woody herbs at Mandalawangi, Suryakencana and Crater Side on the tops Mt. Gede and Mt Pangrango, Cibodas Biosphere Reserve. SEEDLING & HERB 0 50 100 150 200 250 Selliguiea feei Rhododendron retusum Photinia integrifolia Thelymitra javanica Leptospermum flavescens Imperata cylindrica Vaccinium varingiaefolium Gleichenia vulcanica Gaultheria punctata Agrostis infirma Rapanea avenis Rubus lineatus Potentilla indica Hypericum leschenaultii Laurembergia coccinea Anaphalis javanica Lycopodium sp. Gaultheria nummulariodes. Carex verticillata Isachne pangerangensis Tripogon exiguus IMPORTANCE VALUE (%) MANDALAWANGI SURYAKENCANA CRATER SIDE Figure 4. Importance Values of herbs and seedlings at Manda- lawangi, Suryakencana and Crater Side. REINWARDTIA [VOL.12 398 Figure. 5. Species-area curves of seedlings and herbs at Mandalawangi, Suryakencana and the Crater Side on the tops Mt. Gede and Mt. Pangrango, Cibodas Biosphere Reserve. It can be noted from Annex 2 that only one species, Gaultheria nummularioides, occurred throughout the three sites, six species in Mandalawangi and Suryakencana and three species in Suryakencana and the Crater Side, while species with restricted distribution were four only in Mandalawangi, six only in Suryakencana and two only in the Crater Side. Note that in Mandalawangi and Suryakencana, two species of Poaceae (Isachne pangerangensis and Tripogon exiguus) and Cyperaceae (Carex verticillata) were prevalent in this stratum, while in the Crater Side a species of fern, Selliguiea feei, was dominant (Figure 4). The association of Isachne pange- rangensis and Tripogon exiguous is positively correlated in Mandalawangi (r = 0.597) but the correlation is negative in Suryakencana (r = – 0.610), which might be attributed to different habitat conditions. Outside the plots we recorded five additional grass species (Agrostis infirma, Deyeuxia australis, Vulpia bromoides, Isachne globosa and Poa annua). Van Steenis et al. (1972, 2006), however, recorded only five species in the Mt,. Gede-Pangrango area. Vulpia bromoides is a new record as previously it did not occur in this area (Sumadijaya & Veldkamp, 2009; Sunarno & Rugayah, 1992; Van Steenis et al., 1972, 2006). In Mandalawangi, toward the edge, the community was richer, where grasses Agrostis infirma and Deyeuxia australis formed small patches together with big bunches of sedges. On the southwest side the middle part was covered by a thick carpet of Isachne pangerangensis and under this carpet water was dripping. On the eastern side water was oozing from the soil and during the rainy season it became swampy where grasses and sedges were prevalent and very frequently the moss Sphagnum was dominant and constituted the main contributor in the peat forma- tion (Van Steenis et al., 1972, 2006). As indicated in the method above the communities can be stratified only into two strata, i.e. sapling and shrub stratum and seedling and herb stratum. Beside the stratification, density is the only vegetation parameter that can be best used to indicate the structure of the community. Table 1 shows the density of plants in two strata. In the first stratum the highest densities occurred in the Crater Side, consisting of woody species A. lophanta, G. punctata, L. flavescens and V. vari- ngiaefolium and the densities of these species were low in the seedling and herb layer, where the fern Selliguiea feei was prevalent (Figure 4, Annex 2). The average height of A. javanica was 109 cm in Mandalawangi and 77 cm in Surya- kencana. In the Crater Side, however, it reached up to 143 cm, growing well singly and scattered in fine substrates between volcanic rubbles and boulders, apparently experiencing no competttion from other plants. Seedlings of A. javanica had low density, dominance and importance values in both Mandalawangi and Suryakencana, occupying the 6th rank and 8th rank, respectively (Annex 2). We recorded that the seedling density was 1,304/ha in Mandalawangi, and 15,800/ha in Suryakencana (Table 1). It is implied that A. javanica was hardly regenerating under its own shade indicating the pioneer status of the species. Van Steenis et al. (1972, 2006) noted that A. javanica seedlings grew very slowly on volcanic ash deposits and other substrates around craters. The species was said to be a long-live pioneer and was also capable to invade burned subalpine forests, but later disappeared as the burned forest recovered. This has been shown by the dense lower subalpine forest. On the western slope of the Mt. Gumuruh that was burned in 1914; the burned forest was later invaded by A. javanica and slowly 2009] SADILI et al.: Floristic and structure of subalpine summit on Mt. Gede Pangrango 399 reverting to a dense forest (Docters Van Leeuwen, 1933). Van Steenis et al. (1972, 2006) stated that the slow growth of A. javanica was attributed to poor soil conditions and apparently the weathering and soil formation from volcanic materials were progressing very slowly, while at the same time high rainfall and high porosity of the materials facilitated leaching. Such situations were not favorable for forest regeneration but allowing the growth of A. javanica, which did not require good soils to grow. Fire apparently has been recurrent in the alun- alun and more frequently in recent years in view of the frequent visits of hikers, who camped out conveniently there and often negligently left the campfire burning when they left. Fire seems to be a factor, along with poor soil conditions, that maintain the existence of the A. javanica domina- ted community in the alun-alun. There is a slight indication of tree species aggressively invading the community. This is indicated by low relative density of saplings and seedlings of the tree species in the community such as Albizia lophan- ta, Leptospermum flavescens, Rapanea avenis, Symplocos cochinchinensis, and Vaccinium vari- ngiaefolium (Annexes 1 & 2). Tabel 1. Density (individuals/hectare) of saplings, shrubs, seedlings and herbs within two strata of the communities at Mandalawangi and Suryakencana, where A. javanica was the dominant species in the sapling and shrub stratum. Density (Individuals/ha) Stratum Life form and species Mandalawangi Suryakencana Crater Side 1 Sapling and shrub: (a) all species (b) A. javanica 2,461 1,596 2,960 1,976 5,060 400 2 Seedling and herb: (a) All species (b) A. javanica 2,830,435 1,304 2,721,780 15,800 776,000 0 A. MANDALAWANGI 0 20 40 60 80 100 120 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 QUADRAT % C O V E R Carex verticillata Isachne pangerangensis Tripogon exiggius B. SURYAKENCANA 0 10 20 30 40 50 60 70 80 1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 QUADRAT % C O V E R Tripogon exiguus Isachne pangerangensis Carex verticillata Figure 6. Percentage of cover of Carex verticillata, Isachne pangerangensis and Tripogon exiguus in quadrats at Mandalawangi (A) and Suryakencana (B). REINWARDTIA [VOL.12 400 Based on the Importance Values of sapling- shrub stratum and those on seedling-herb stratum, the Mandalawangi meadow may be designated as the Anaphalis javanica-Isachne pangerangensis community type (Figure 7), the Suryakencana meadow as Anaphalis javanica-Tripogon exiguus community type (Figure 8) and the Crater Side scrub as Vaccinium varingiaefolium-Seliguea feei community type (Figure 9) Using Jaccard’s formula (Mueller-Dombois & Ellenberg, 1974) and the Importance Values, the Index of Similarity (IS) of community types at Mandalawangi and Surayakencana was 95.82% for the sapling stratum and 75.23% for seedling stratum, res- pectively. The similarities between the Crater Side community type and the community types at Mandalawangi and Suryakencana at the sapling stratum are high, with IS of 69.60% and 78.93%, respectively, while at the seedling stratum was very low, with IS of 2.11% and 9.72%, respectively. These community types are simi- lar to those at the alun-alun in Mt. Papandayan (Van Steenis, 1930, 1932), and Mt. Ceremai (Lam, 1925), West Java and Mt. Sumbing and Mt. Sindoro (Docters van Leeuwen, 1932) Central Java, although A. javanica occurred less abundantly. Floristically at generic level and structurally these community types are comparable to those in the subalpine regions in various mountains in Indonesia, such as in Papua (Hope, 1976; Johns et al., 2007), while communi- ty type dominated by Vaccinium varingiaefolium is common as pioneer vegetation in the volcanic mountains elsewhere, particularly in Java, e.g. in Mt. Guntur (Van Der Pijl, 1938), Mt. Ceremai, (Lam, 1925), Papandayan (Van Steenis, 1930, 1932, 1935; Van Steenis et al., 1972, 2006; Docters Van Leeuwen, 1932) and Mt. Tangkuban Perahu (Wiljes-Hissink, 1953). Another important and widespread species was Carex verticillata (Figure 6 & Annex 2), whose cover tends to increase and it takes over the dominance when I. pangerangensis or Tripogon exiguus was low in cover or absent in the community (R cover of –0.31 and 0.19, respectively, in Mandalawangi and –0.24 and – 0.19, respectively, in Suryakencana) as indicated also in Figure 4. In Suryakencana, Agrostis infirma and Gleichenia vulcanica may become important also in places. The presence of such correlations reflects the pattern in vegetation and is usually related to habitat factors, yet to be investigated thoroughly. The results of the present investigation show that the meadows in Mandalawangi and Suryakencana were similar, but they were relatively poor in species, confirming the earlier records. It has been assumed that poor soils and fires have maintained the perpetual existence of A. javanica in the meadows. 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List of species of saplings, shrubs and woody herbs with their values on Relative Dominance (RDo), Relative Density (RD), Relative Frequency (RF) and Importance Value (IV) at the Mandalawangi, Suryakencana and the Crater Side at the tops of Mt. Gede-Pangrango, Cibodas Biosphere Reserve. MANDALAWANGI SURYAKENCANA CRATER SIDE NO. SPECIES FAMILY RDo RD RF IV RDo RD RF IV RDo RD RF IV 1 Anaphalis javanica Asteraceae 69.0 70.0 48.0 187.0 82.8 66.8 44.8 194.3 1.0 1.0 6.6 8.5 2 Vaccinium varingiaefolium Ericaceae 6.3 2.5 8.1 16.8 4.7 7.6 13.3 25.6 72.0 78.9 32.8 183.7 3 Leptospermum flavescens Myrtaceae 5.0 11.1 10.4 26.5 1.8 3.6 8.1 13.5 1.0 0.2 1.6 2.9 4 Gaultheria punctata Ericaceae 0.8 1.1 4.1 6.0 7.6 15.9 19.5 43.0 8.6 5.0 16.4 30.0 5 Symplocos cochinchinensis Symplocaceae 3.4 0.8 5.2 9.4 0.4 1.1 2.6 4.1 0.9 0.2 1.6 2.8 6 Rubus lineatus Rosaceae 0.4 2.1 2.9 5.4 0.1 0.3 0.7 1.1 0.7 0.2 1.6 2.6 7 Eurya glabra Theaceae 2.9 0.3 1.7 4.9 0.2 0.2 0.7 1.1 0.6 1.0 4.9 6.5 8 Lonicera javanica Caprifoliaaceae 2.1 1.4 4.6 8.1 0.8 1.8 3.3 5.9 9 Photinia integrifolia Rosaceae 0.8 1.3 5.2 7.4 1.1 1.8 3.8 6.6 10 Rapanea avenis Myrsinaceae 1.7 8.0 3.5 13.1 0.1 0.3 0.5 0.9 11 Vaccinium laurifolium Ericaceae 1.3 0.7 2.9 4.9 0.3 0.3 1.0 1.6 12 Rubus sp. Rosaceae 0.0 0.1 0.2 0.3 13 Dicrochepala chrysantemifolia Asteraceae 0.1 0.1 1.0 1.1 14 Albizia lophanta Fabaceae 0.0 0.1 0.5 0.7 3.1 1.4 6.6 11.3 15 Rhododenron retusum Ericaceae 11.7 11.3 24.6 47.7 16 Scheflera lucescens Araliaceae 0.6 0.7 3.3 4.6 17 Malus domestica Rosaceae 5.4 0.1 0.6 6.1 18 Hypericum leschenaultii Ericaceae 0.8 0.8 2.9 4.6 Total 100 100 100 300 100 100 100 300 100 100 100 300 2009] SADILI et al.: Floristic and structure of subalpine summit on Mt. Gede Pangrango 403 Annex 2. List of species of seedlings and herbs with percentages of their Relative Dominance (RDo), Relative Density (RD), Relative Frequency (RF) and Importance Value (IV) at Mandalawangi, Suryakencana and the Crater Side at the top of Mt. Gede- Pangrango No SPECIES FAMILY MANDALAWANGI SURYAKENCANA CRATER SIDE RDo RD RF IV RDo RD RF IV RDo RD RF IV 1 Gaultheria nummulariodes. Ericaceae 0.4 0.5 16.53 7.43 0.79 0.63 5.88 7.30 0.2 0.3 7.69 8.2 2 Hypericum leschenaultii Hypericaceac 0.1 0.2 0.83 1.13 3 Laurembergia coccinea Haloragaceae 1.7 0.1 4.96 6.76 4 Rubus lineatus Rosaceae 0.4 0.2 0.83 1.43 5 Rapanea avenis Myrsinaceae 0.2 0.4 0.83 1.43 6 Tripogon exiguus Poaceae 38.5 28 18.18 84.68 54.7 34.69 12.25 101.60 7 Isachne pangerangensis Poaceae 30.3 48 19.01 97.31 7.05 30.13 12.25 49.43 8 Carex verticillata Cyperaceae 18.7 19 18.18 5.88 18.3 24.51 12.25 55.08 9 Anaphalis javanica Asteraceae 2.6 1.9 11.6 6.07 1.53 0.58 8.09 10.20 10 Potentilla indica Rosaceae 0.2 0.5 0.83 1.53 1.17 0.2 3.68 5.05 11 Lycopodium sp. Lycopodiaceae 7 1.2 8.26 16.46 0.93 1.74 9.80 12.47 12 Thelymitra javanica Orchidaceae 0.47 0.02 0.25 0.74 13 Gleichenia vulcanica Gleicheniaceae 2.4 2.12 7.35 11.87 14 Imperata cylindrica Poaceae 0.74 0.11 2.94 3.79 15 Leptospermum flavescens Myrtaceae 0.35 0.04 0.49 0.88 16 Agrostis infirma Poaceae 5.92 4.08 12.25 22.25 17 Photinia integrifolia Rosaceae 0.47 0.01 0.25 0.73 18 Vaccinium varingiaefolium Ericaceae 2.56 0.13 3.43 6.12 17.4 9 23.08 49.5 19 Gaultheria punctata Ericaceae 2.6 1 8.58 12.18 1.1 0.3 7.69 9.1 20 Rhododendron retusum Ericaceae 0.04 0.01 0.25 0.30 2.5 0.8 23.08 26.4 21 Selliguiea feei Polypodiaceae 78.7 89.7 8.46 206.9 TOTAL 100 100 100 300 100 100 100 300 100 100 100 300 Figure 7. The subalpine Anaphalis javanica-Isachne pangerangensis community type in Alun-alun Mandalawangi at the Mt. Pangrango summit, surounded by subalpine Leptospermum flavescens forest. REINWARDTIA [VOL.12 404 Figure 8. An overview of the subalpine Anaphalis javanica-Tripogon exiguous community type at Alun-alun Suryakencana in the Mt. Gedeh summit, complex, surrounded by subalpine forest dominated by Vaccinium varingiaefolium and Leptospermum flavescens or in places by Albizia lophanta Figure 9. The subalpine Vaccinium varingiaefolium-Seliguea feei community type, surrounded by subalpine Albizia lophanta forest at the Crater Side in the Mt. Gede summit complex.